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This appears to be
yet another case of pertinent taxon exclusion. Today’s fossorial digger has several universally acknowledged xenarthran (edentate) traits. For reasons unknown it was not tested against another fossorial xenarthran, Peltephilus. Rather the authors compared their digger to an arboreal sloth, Bradypus, among several other taxa, including distinctly different anteaters and armadillos.

ºªº Figure 1. Scapula of Fruitafossor compared to several candidate sisters. Luo and Wible made things a bit more difficult by presenting left and right scapulae. In frame 2 they are all left scapulae for ready comparison. There is no doubt that the Fruitafossor scapula was illustrated to look more like that of Ornithorhynchus. Unfortunately the photo data (Fig. 2) does not clearly support that shape. That shape is so important, it needed to be better documented.

Luo and Wible 2005brought us a small, mostly articulated, rather crushed and incomplete Late Jurassic mammal with simple blunt teeth and digging forelimbs. Fruitafossor windscheffeli (Figs. 1–6) is best represented by a CT scan (Figs. 2–4) and original drawings (Figs. 5, 6) created by the Luo and Wible team.

Figure 2. Fruitafossor in situ from Digimorph.org and used with permission and here colorized to an uncertain extent. All those little white dots could be scattered osteoderms.

The original analysis
nested Fruitafossor between extremely tiny Hadrocodium + Shuotherium and Gobiconodon in a tree topology that does not resemble the topology of the large reptile tree (LRT, 1048 taxa). The authors noted Fruitafossor is “not a eutherian, let alone a xenarthran”despite noting Fruitafossor had tubular molars and xenarthran intervertebral articulations, traits otherwise found only in xenarthrans.

Figure 3. Same specimen from Digimorph.org and rotated to show the teeth better. See figure 3 for a closeup. All those little white dots could be scattered osteoderms. Some of this flatness is due to crushing. Some of it is due to this being a wider than deep armored mammal.

Wider than deep
Yes, the Fruitafossor specimen is crushed, but what is shown here indicates a low, wide mammal just getting some armor in the Late Jurassic. And we all know why armor might have been helpful! And this may explain the lateral sprawl of the forelimbs and giant wide humerus, another atavism!

Figure 4. Closeup of figure 2 showing maxilla and dentary in occlusion. If there is a convex ventral dentary it must be imagined because it is not preserved.

Figure 5. The xenarthran, Peltephilus, compared to Fruitafossor, not to scale, but to similar jaw lengths. Note the drawing from Luo and Wible does not exactly match what one can see in the photo (Fig. 3). This data needs to be clear and it is not.

Luo and Wible comparedFruitafossor to the arboreal and extant Bradypus, but not to the fossorial and extinct Peltephilius (Fig. 5). I would consider that a mistake or an oversight that here overturns their hypothesis of a relationship of Fruitafossor to basalmost mammals.

Figure 6. Several drawings from Luo and Wible that one must trust for accuracy. The verification data is too fuzzy to validate. As in other xenarthrans, the ilia actually form a pair of horizontal plates on either side of the long fused and eroded sacrals. Four fingers is a trait shared with Peltephilus. Imagine that rib cage wider and not so deep.

Scattered osteoderms(Fig. 3) were not mentioned in the text. That’s one more trait shared with the armored xenarthran, Peltephilus. This overlooked relationship of derived xenarthrans moves them into the Jurassic, an era they have never been in before in phylogenetic and chronological analyses. Here placental arboreal and fossorial mammals (prior to condylarths) shared time and space with dinosaurs during the Jurassic and Cretaceous, but have, so far, been underrepresented in the fossil record. That’s changing with re-examination of the data applied to a larger gamut taxon list.

Although the illustrated scapula
(Fig. 1) looks like that of an egg-laying mammal, I will wait for better data to validate that illustration. In the meantime, Fruitafossor has blunt, tubular molars and xenarthran vertebral articulations (among many other xenarthran traits) because it is a xenarthran, not an egg-layer with convergent traits.

PrefaceI’ve been wondering about the traditional nesting of Multituberculata and kin outside of the Mammalia for years. All have a dentary jaw joint, but some have post-dentary bones. given the opportunity multituberculates nest with rodents and plesiadapiformes in the large reptile tree (LRT, 1047 taxa). No other pre-mammals resemble them. Traditionally Haramiyava(Fig. 1) has been considered a pre-mammal link to Haramiyida + Multituberculata. In the LRT Haramiyava nests with the mammaliaforms Brasilodon, Sinoconodon and Therioherpeton– far from any other taxa considered Haramiyida + Multituberculata currently and provisionally nesting deep within the Mammalia.

Figure 1. Haramiyavia reconstructed and restored. Missing parts are ghosted. The fourth maxillary tooth appears to be a small canine. The post-dentary bones are imagined from Vilevolodon (figure 4).

Vilevolodon diplomylos(Luo et al. 2017; Jurassic, 160 mya; BMNH2942A, B; Figs. 2-4) was originally considered a stem mammal (= mammaliaform), a eleutherodontid in the clade Haramiyida AND it had clearly defined gliding membranes (Fig. 2). By contrast the LRT nests Vilevolodon with the Late Jurassic para-rodent Shenshou and the extant rodents, Rattus and Mus, not far from members of the Multituberculata.

The ear problem in Jurassic rodents
Luo et al. report, “a mandibular middle ear with a unique character combination previously unknown in mammaliaforms.”Pre-mammals have post-dentary bones (articular, angular, surangular). Therian mammals shrink and migrate those bones to the base of the skull where they become middle ear bones with new names (malleus, incus, ectotympanic). The stapes remains the stapes in all tetrapods. So what is happening with Vilevolodon and its sisters? Why don’t the pre-mammal post-dentary bones define it as a pre-mammal? After all, that’s the current paradigm.

Figure 3. There is no doubt that Vilevolodon has pre-mammal type post-dentary bones. There is also no doubt that the dentary formed the main jaw joint with the squamosal. How does one reconcile both sets of traits? In the LRT Vilevolodon nests with rodents. This appears to be a mammal with an atavism, a reversal. These elements simply stopped developing as in other mammals.

Mammals are defined by
the evolution and migration of their posterior jaw bones into middle ear bones with a jaw joint switch from quadrate/articular to dentary/squamosal. Multituberculates and haramiyids appear to bend or break that rule because they have cynodont-like posterior jaw bones, not tiny middle ear bones, and yet otherwise they nest with rodents and plesiadapiformes. This is one reason why you don’t want to pull a Larry Martin with post-dentary bones. You want to nest a taxon based on a long list of traits, not just one, two or a dozen.

The massive jaw jointMammals, such as Vilevolodon, with atavistic post-dentary bones also have a massive jaw joint with a long articulating surface on the dentary contacting the squamosal. All mammals have such a jaw joint. Pre-mammals don’t. While Vilevolodon has a large dentary/squamosal jaw joint, the post-dentary articular, still contacts the quadrate. It’s clearly not the main jaw joint.

Filan 1991
traced the development of post-dentary bones in embryonic Monodelphis specimens. She reported, “Neonates of Monodelphis possess neither mammalian (dentarysquamosal) nor reptilian (quadrate-articular) jaw articulations, nor does the contact between the incus and crista parotica offer a joint surface. Elasticity in Meckel’s cartilage allows minimal deflection of the lower jaw.”After all, those neonates are just sucking milk, not biting, and the embryos don’t even do that. Does that make neonates like this not mammals? No. The evidence indicates that in multituberculates and haramiyds the embryological transformation of posterior jaw bones stopped before development transformed them into middle ear bones. This is an atavism, a phylogenetic reversal. The timing of development changed. In the case of Vilevolodon, the middle ear bones stop evolving during embryological development and the post-dentary bones they would have evolved from continue to appear in adults. What was a rare mutation probably spread throughout an isolated population. Perhaps this had something to do with the increase in size of the dentary jaw joint.

Haramiyavia and the Haramiyida clade
Seems at this point that only Haramiyavia is a haramiyid, unless Brasilodon is one as well. Members traditionally assigned to the cladeEleutherodontidae also nest in various locations in the LRT, not all in one clade.

Meng et al. 2017 report,“Stem mammaliaforms are morphologically disparate and ecologically diverse in their own right, and they developed versatile locomotor modes that include arboreal, semiaquatic, and subterranean specializations, which are all distinct from generalized mammaliaforms.” Unfortunately, the LRT nests a long list of mammaliaforms at various nodes within the Mammalia. They are not from a single diverse clade.

Contra Meng et al. 2017
the LRT reduces the niches and body shapes of stem mammals down to a few small, generalized taxa like Sinoconodon and Megazostrodon. Derived taxa nest at derived nodes.

The LRT nests rodents close to Plesidapiformes,including the extant aye-aye, Daubentonia as first reported here.So it comes as no surprise when Luo et al. report, “Eleutherodontids show a marked similarity to the primate Daubentonia in the ventrally bent rostrum and deep mandible, and both features are interpreted to be reinforcement for incisor gnawing.” That’s the case only with Vilevolodon this time. Others may be by convergence.

Molars
The jaw joint of the rodent allows for rostral-caudal and dorsal-ventral motion of the jaws. Luo et al. report, in Villevolodon it is not possible for the mandible to move posteriorly or horizontally, but their images show a continuous anteroposterior trough/furrow in the three molars, though not to the extent seen in sister taxon Shenshou. Molars with a long and continuous trough for rostral-caudal grinding appear by convergence in several reptile/mammal clades.

Incisor replacement
Luo et al. report, “Incisor replacement is prolonged until well after molars are fully erupted, a timing pattern unique to most other mammaliaforms.“ In rodents incisors never stop growing. The growth pattern in Vilevolodon may be the first step toward that. Not sure why Luo et al. are missing all these strong rodent clues.

Gliding?Meng et al. 2017 note: “They [Vilevolodon and kin] are the most primitive known gliders in mammal evolution, evolving approximately 100 million years before the earliest known therian gliders.” Earlier, with the appearance of the stem pangolin, Zhangheotherium at the start of the Cretaceous, the ghost lineage for primates, flying lemurs and bats was also set to that time or earlier. Before the advent of flying birds, but after the advent of predatory theropods, many mammals had evidently taken to the trees. And one way to get from tree to tree without descending to the dangerous turf is to jump, glide and fly. I predict we’ll find the big-handed ancestors of bats in Jurassic and Cretaceous strata someday. They are already volant shortly after the K-T extinction event.

Hearing in Vilevolodon
With the reappearance of post-dentary bones in taxa like Vilevolodon, the auditory acuity that was more highly developed in its ancestors must have suffered a setback. By the evidence provided, the massive jaw joint must have been more important for its survival.

Figure 4. Multituberculate Kryobaatar mandible in lateral and medial views. Here post-dentary bones are absent here. The malleus (quadrate) and ectotympanic are on the skull.

Getting back to the purported patagium of Maiopatagiumwhich we looked at yesterday. It is not apparent and the authors do not describe it. Rather, Meng et al. 2017 sidestep this by reporting, “Furthermore, we report a second eleutherodont specimen (BMNH2942) preserved with a halo of carbonized fur and patagial membranes, similar to those of Maiopatagium.”The patagial taxon remains unnamed in the Maiopatagium paper (Meng et al. 2017), but is named in a second paper appearing on the same day. It is today’s subject, Vilevolodon (Fig. 1)

Figure 1. Yanoconodon fossil in situ. See the skull in closeup in figure 2. The published tracing is distorted here to match the underlying photo.

Wikipedia reports,“Yanoconodon was a small mammal, barely 5 inches (13 centimetres) long. It had a sprawling posture, Yanoconodon was a Eutriconodont, a group composing most taxa once classified as “triconodonts” which lived during the time of the dinosaurs. These were a highly ecologically diverse group, including large sized taxa such as Repenomamus that were able to eat small dinosaurs, the arboreal Jeholodens, the aerial volaticotherines and the spined Spinolestes. Yanoconodon is inferred to be a generalized terrestrial mammal, capable of multiple forms of locomotion.

Figure 2. Yanoconodon is exposed in ventral view. Even so, if you employ DGS, even on a fuzzy photo, you can put together a reconstruction that shares several traits with Repenomamus.

Mammal-like reptiles?Wikipedia also reports, “The Yanoconodon holotype is so well preserved that scientists were able to examine tiny bones of the middle ear. These are of particular interest because of their “transitional” state: Yanoconodon has fundamentally modern middle ear bones, but these are still attached to the jaw by an ossified Meckel’s cartilage.This is a feature retained from earlier stem mammals, and illustrates the transition from a basal tetrapod jaw and ear, to a mammalian one in which the middle ear bones are fully separate from the jaw. Despite this feature Yanoconodon is a true mammal. It is thought that the feature was retained during early embryo development,[4] whereas it is lost in most other mammal groups. The intermediate anatomy of the middle ear of Yanocodon is said to be a “Rosetta Stone”[5] of mammalian middle ear evolution.”

In the large reptile tree (LRT, 1037 taxa) Yanoconodon, Repenomamus,Jeholodens and Spinolestes are not mammals, but very close to the base of the Mammalia. Both clades share Pachygenelus as last common ancestor. So that means the ‘transitional state’mentioned above is indeed outside the Mammalia. Other paleontologists consider this list of taxa to be mammals, but here the mammal-like traits they had were developed in parallel and not quite to mammal standards.

Figure 4. Repenomamus reconstructed using DGS methods. The manus and feet are loose figments at present. Despite its predatory nature, note the reduction in canines, a clade trait.

The skull of Yanoconodon(Fig. 2) can be largely, but not completely, reconstructed based on the visible bones. The skull is low and wide and without the typical constriction anterior to the jugals. The anterior teeth are large and spike-like while the posterior teeth are molariform. Large teeth typically require deep roots and deep bones to house those roots. The mandibles are as long as the skull. The small orbits are far forward on the skull and the temporal fenestra are correspondingly large.

Figure 2. The origin and radiation of stem mammals and crown mammals. Compare the LRT tree (above) to a recent cladogram by Close et al. 2015.

With the new data on Yanocondonseveral taxa within the LRT shifted places, but not far and still within the derived Cynodontia. Something about the Mammalia helped them survive several extinction events that the derived Tritylodontia (= Pseudomammalia) succumbed to. Pseudomammalia LOOK like mammals, but are not mammals. They continued to exist into the Early Cretaceous and some, like Repenomamus, were quite large.

Figure 1. Vintana as originally illustrated. I added colors to certain bones. Note the high angle of the ventral maxilla and the deep premaxilla. Lateral view reduced to scale with other views.

Earlier we looked at Vintana (Fig. 1, Krause et al. 2014a, b). To Krause et al. Vintana represented the first specimen in the clades Allotheria and Gondwanatheria to be known from more than teeth and minimal skull material.

Despite a paper in Nature
and a memoir of 222 pages in the Journal of Vertebrate Paleontology; despite CT scans and firsthand examination with electron microscopes; despite being examined and described by many of the biggest name and heavy hitters in paleontology… Krause et al. never understood that Vintana was just a derived wombat, evidently due to taxon exclusion problems.

Figure 2. Interatherium does not nest with notoungulates or other purported interotheres. Rather cat-sized Interatherium nests with wombats,with Vintana, between Vombatus and the giant Toxodon

The large reptile tree now includes
1005 taxa, all candidates for sisterhood with every added taxon. Despite the large gamut of 74 taxa employed by Krause et al. they did not include the best candidates for Vintana sisterhood. Perhaps the fault lies in the reliance of prior studies and paradigms. Perhaps the fault lies in the over reliance by Krause et al. and other mammal workers, on dental traits. Perhaps the fault lies in the absence of pertinent sisters to the above-named taxa, including Interatheriium for Vintana.

In any caseVintana does not stand alone as the only taxon in its clade represented by skull material. Based on its sisterhood with Interatherium, we have pretty good idea what its mandibles and post-crania looked like. Yes, Vintana is weird. But Interatherium is also weird in the same way, just not as weird.

It’s an early Eocene pig
according to the large reptile tree (LRT, 1003 taxa). A large gamut minimizes inclusion set bias and gains greater authority with every added taxon. It also reduces the average phylogenetic distance between taxa, all of which are species and individuals, not suprageneric taxa.

Figure 1. Danjiangia nests with the extant pig, Sus, in the LRT. Note the very low naris and nasal. The lost skull could have been elevated, as imagined here after phylogenetic analysis.

Danjiangia pingi(Wang 1995; early Eocene) was originally described as a basal chalicothere with brontothere traits. Then Hooker and Dashzeveg (2003) nested it as a basal brontothere (without including any other brontotheres). Mihlbacher 2008 and others used it as an outgroup to the brontotheres. The posterior skull is not known, but note the rise over the orbits suggesting a tall cranium, as in Sus (Fig. 2). Also note the very low naris below the low nasals. Usually you don’t see nasals so low, and perhaps that is due to taphonomic shifting.

Figure 2. Skull of the extant pig, Sus in several views. Note the elevated cranium and squamosal.

The long fused dentary
of Danjiangia is a trait also shared with pigs and other taxa, like chalicotheres, by convergence.

Figure 3. Skeleton of Sus, the pig. It provides good clues as to the missing postcranial skeleton of Danjiangia.

Sus the pig
(Fig. 3) provides good clues as to the missing postcranial skeleton of its sister taxon, Danjiangia. The other model for post-cranial details is the basal artiodactyl, Cainotherium(Fig. 4).

Fig. 4. Cainotherium nests at the base of the Artiodactyla or even-toed ungulates. I wonder if it had five fingers, even if vestiges, given that Ancodus, a derived artiodactyl, retains five fingers.

Why was the pig connection missed by others?
For the same reason that modern workers continue to include pterosaurs with archosaurs. It’s a tradition. Nobody wants to do the extra work of testing other candidate taxa. Nobody wants to acknowledge contrarian studies. Paleontology tends to run very slowly as we learned earlier here. Hail, hail the status quo!

Figure 5. Subset of the LRT focusing on ungulates, which split into three clades here. Note the nesting of Sus together with Danjiangia.

ReferencesBeard KC 1998. East of Eden: Asia as an important center of taxonomic origination in mammalian evolution; pp. 5–39 in Beard and Dawson (eds.), Dawn of the Age of Mammals in Asia. Bulletin of Carnegie Museum of Natural History 34.Mihlbachler MC 2004. Phylogenetic systematics of the Brontotheriidae (Mammalia, Perissodactyla). PhD dissertation. Columbia University. p. 757.Mihlbachler MC 2008. Species taxonomy, phylogeny and biogeography of teh Brontotheriidae (Mammalia: Perissodactyla). Bulletin of the American Museum of Natural History 311:475pp.Wang Y 1995. A new primitive chalicothere (Perissodactyla, Mammalia) from the early Eocene of Hubei, China. Vertebrata Palasiatica 33: 138–159.

In 1873
O. C. Marsh 1874) found an extinct Eocene (50.3 to 40.4 Ma) mammal “of great interest. The lower molar teeth, all essentially alike, and inserted in deep sockets” were the most striking feature. He named it Stylodon mirus(Figs. 1,2). All the teeth grew with “persistent pulps” and had a thin layer of enamel. The specimen was considered close to Toxodonwith some edentate affinities (Marsh 1897). Stylinodon was placed under the family Stylinodontidae and the order Tillodontia. According to Schoch 1986 (first issue of JVP!) its ancestors were like Onychodectes.

Stylinodon mirus(Marsh 1874; middle Eocene, 45 mya; Figs. 1-2) was originally considered a taeniodont, perhaps derived from Onychodectes. Here it nests with Mustela, the living European mink, among the Carnivora. There were twice as many molars (4), each with a single root, as in the two double rooted molars of the mink. Large claws and certain forelimb traits indicate that Stylinodon was a digger, not a cursor.

The present nestingof Stylinodon mirus(YPM VP 011095, Marsh 1874; Figs. 1, 2) in the Carnivora occurred when I realized it was a poor fit at the base of the Condylarthra/Paenungulata, despite its herbivorous dentition and tusk-like teeth (canines, not incisors).

Figure 1. Stylinodon skull. Note the transverse premaxilla, a trait of the Carnivora.

Distinct from condylarthsStylinodon has a transverse premaxilla, essentially invisible in lateral view. The lower canine is the anteriormost tooth on the dentary. These traits are shared with other members of the Carnivora. In the present taxon list Stylinodon shares more traits with Mustela, the European mink (Fig. 1) despite the loss of molar cusps and increase in size. They both were diggers. Together they nest with Phoca, the seal, and Palaeosinopa, the amphibious piscivore, all derived from a sister to Procyon, the omnivorous raccoon (Fig. 2).

Figure 3. Mustela the European mink is an extant relative to Stylinodon.

Mustela lutreola(Linneaus 1761; extant European mink; up to 43cm in length) is a fast and agile animal related to weasels and polecats. Mustela lives in a burrow, but it also swims and dives skilfully. It is able to run along stream beds, and stay underwater for one to two minutes. Mustela is derived from a sister to Phoca and other seals, all derived from a sister to Procyon. With this close relationship, Stylinodon (Fig. 2 a giant weasel with simple teeth.

Schoch and Lucas 1981and Schoch 1983 considered Stylinodon and kin derived from a sister to the long-legged basal condylarth, Onychodectes. The large reptile tree (LRT, Fig. 2) does not support that nesting. Onychodectes has a long premaxilla lacking in taeniodonts.

Figure 4. Subset of the LRT showing the Carnivora nesting at the base of the Eutheria (placental mammals).

Schoch and Lucas 1981determined that Stylinodon had two upper incisors (one lower), a giant canine, four premolars and three molars, as in Onychodectes. That may be so, but the premolars and molars look alike.

Wortmania (Hay 1899, Williamson and Brusatte 2013; above) and Psittacotherium(Cope 1862; below) are related to Stylinodon. All are among the largest taxa in the early post-Cretaceous, derived from smaller weael-like basal mammals in the Cretaceous.

Figure 6. Psittacotherium in various views. Overall it is elongated to more closely match related taxa.

It is rare but not unheard offor members of the Carnivora to become omnivores and herbivores. Think of the giant panda and certain viverrids. Now the stylinodontid taeniodonts join their ranks.

ReferencesLinneaus C von 1761. xxxMarsh OC 1874. Notice of new Tertiary mammals 3. American Journal of Science. (3) 7i: 531-534.|Marsh OC 1897. The Stylinodontia, a suborder of Eocene Edentates. The American Journal of Science Series 4 Vol. 3:137-146.Rook DL and Hunter JP 2013. Rooting Around the Eutherian Family Tree: the Origin and Relations of the Taeniodonta. Journal of Mammalian Evolution: 1–17.Schoch RM and Lucas SG 1981. The systematics of Stylinodon, an Eocene Taeniodont (Mammalia) from western North America. Journal of Vertebrate Paleontology 1(2):175-183.Schoch RM 1983. Systematics, functional morphology and macroevolution of the extinct mammalian order Taeniodonta. Peabody Museum of Natural History Bulletin 42: 307pp. 60 figs. 65 pls.

As usualI had second hand (academic papers and figures) rather than firsthand access to the specimens. It doesn’t matter how good your players are if you don’t show up on the right field at the proper hour. Here you’ll see, once again, how excluding the actual sister to an enigma taxon is the major problem, solvable by second-hand phylogenetic analysis in a large gamut study, the large reptile tree (LRT) that minimizes the problem of taxon exclusion.

Figure 1. Necrolestes skull. Note the scale bar problems. DGS colors the bones here. The lacrimal and infraorbital are enlarged here, providing a large opening for large facial nerves. Note the larger lower incisors as compared to the drawing above.

Necrolestes patagonensis (Ameghino 1891; early Miocene, 16mya; Fig. 1; YPM PU 15065, 15384, and 15699) has been argued about for over a hundred years. Originally (Ameghino 1891) it was described as the only known extinct placental “insectivore” from South America and allied to Chrysochloris (Fig. 2), the extant golden mole.

Well done Ameghino!

Unfortunately, as time went on…
Saban 1954 considered Necrolestes a palaeanodont (Ernanodonwas previously considered one). Patterson 1958 considered it a borhyaenoid metatherian. Asher et al. 2007 looked at several candidates and could not make a firm conclusion. Ladevèze et al. 2008 supported metatherian affinities. Goin et al. 2008 also could not be specific with regard to a closest known sister taxon.

The latest paper on the subjectRougier et al. 2012 reported, “earlier studies leaned toward placental affinities and more recent ones endorsed either therian or specifically metatherian relationships.” Ultimately they nested Necrolestes with Cronopio (Fig. 4) which they considered a non-therian mammal. That is correct. They considered an earlier Van Valen 1988 statement inspired, “…the enigmatic Miocene genus Necrolestes, usually thought to be a marsupial, is [conceivably] a late surviving Gondwantherian pantothere.” That is incorrect.

Figure 2. Chrysochloris skull lateral view. Note the many similarities to Necrolestes, including a ventral naris, dorsally expanded bulla, and similar shapes for the other bones. Note the orbit is very tiny in this burrowing taxon. I don’t see an infraorbital foramen. here, distinct from Necrolestes.

Asher et al. 2007 report,“Characters that support [Necrolestes] status as a therian mammal include a coiled cochlear housing of the inner ear. Necrolestes shows similarities to eutherian mammals, such as small incisive foramina and possibly three molars.Consistent with its status as a metatherian is the presence of five upper incisors, transverse canal foramina, and a broad proximal fibula. However, we cannot confirm other characters claimed by previous researchers as evidence for affinity with marsupial or nonplacental mammals, such as the presence of an inflected mandibular angle and epipubic bones.”

Asher et al. report,“The idea that [Necrolestes] is related to golden moles was favored in the first two publications describing its anatomy (Ameghino, 1891; Scott, 1905). We do not believe Patterson’s contention that the status of Necrolestes as a marsupial is ‘‘virtually assured’’. We admit that the list of possible taxonomic affiliations for this animal still remains long.”

Figure 3. The Golden Mole (Chrysochloris asiaticus) nests with the tree shrew and elephant shrew in the large reptile tree, not the common mole. Image copyright Digimorph.org and used with permission.

The large reptile tree
(920 taxa) tested Necrolestes against a wide gamut of mammal candidates and nested it securely with the golden mole, Chrysochloris. To shift Necrolestes next to Cronopio adds 22 steps.

Distinct from sister taxaNecrolestes had five upper incisors and four lowers. That is closer to the primitive numbers for mammals and two more than in Chrysochloris. The molars are also primitive in having fewer cusps, but that also happens in whales and armadillos… and golden moles… with their simplified zalambdodont teeth… so let’s focus on other traits. Dental traits are plastic and can lead analysis astray.

Rougier et al. report,“the first upper and lower premolars are double rooted and the following five molariform elements are single rooted, a condition shared only with the recently described meridiolestidan mammal Cronopio.”Convergent dental traits might be leading these workers so far afield the neglected to add Chrysochloris to their analysis, which seems odd and dangerous based on the long list of shared traits and overall similarity, not by convergence.

Figure 4. Cronopio nests between Juramaia and Didelphis + Ukhaatherium in the LRT. Rogier et al. nest this taxon with Necrolestes, contra the LRT. This taxon has an anterior naris, not a ventral one.

Rougier et al. gave us straw dogs
when they compared the basicrania of several sister candidates, but NOT that of Chrysochloris, to that of Necrolestes. Here I add a basicranium Rougier et al. chose to not show. Chrysochloris more closely matches the morphology of Necrolestes than any of the other three candidates. I don’t see Chrysochloris listed in the Supplemental Information for Rougier et al. which appears to test non-placental mammals only. So this is what I mean by another case of taxon exclusion. Ameghino (1891) got it right originally. Rougier Wible, Beck and Apesteguía 2012, for some reason, dropped the ball.

Figure 3. Necrolestes basicrania compared to three candidates by Rougier 2012. Here I add the basicranium for Chrysochloris for comparison and it’s a better match. The blue element is the posterior mandible, which is not shown on the Rougier et al. drawings. Not how the lower (posterior) element curls over the basicranial element in only two candidates here. This is a placental trait. The LRT uses no petrosal traits, but image speaks for itself. Excluding the actual sister taxon was done for reasons unknown in this flawed study.

Deleting Chrysochloris from the LRTnests Necrolestes with the remaining basal Glires, but resolution is lost. Not sure why, but Necrolestes has a history (see above) of being a confusing taxon when not nested with Chrysochloris.

Deleting all placentals from the LRT,except Necrolestes, nests it between Didelphis and Asioryctes a node apart from Cronopio. So taxon exclusion doesn’t recover what Rougier et al. recovered.

Now that we have golden moles in Africa and South Americathis is evidence that golden moles first appeared before those continents split apart 118 to 115 mya, long before the end of the Cretaceous. Video link here. Naish reports, “Golden moles and tenrecs appear to be close relatives, forming a clade usually termed Afrosoricida Stanhope et al., 1998 (though this is essentially synonymous with Tenrecoidea McDowell, 1958, see Asher (2001)“. That relationship is not supported by the LRT. Golden moles probably first appeared in the Early Jurassic, given that other Glires, multituberculates, split from rodents about the same time and are found as early as Middle Jurassic strata.

Rougier et al. tested earlier studies and found them flawed
Similarly, I tested Rougier et al. and found it flawed. Perhaps someday someone will likewise test this test and present additional insight into this former enigma taxon.